Applied Animal Behaviour Science 78 (2002) 263±290

Equid play ethogram

Sue M. McDonnell*, Amy Poulin
Equine Behavior Laboratory, New Bolton Center, University of Pennsylvania School of Veterinary Medicine,
382 West Street Road, Kennett Square, PA 19348, USA

Abstract

An ethogram of play behavior among equids was developed. Several key English-language studies
on equids were reviewed to derive a preliminary inventory of speci®c behaviors to be included in the
ethogram. Our primary observations were based on a herd of semi-feral Shetland-type ponies kept at
New Bolton Center, University of Pennsylvania School of Veterinary Medicine, Kennett Square, PA.
Greater than 100 h of direct observation and photo-documentation focused speci®cally on play in
order to identify play behaviors to be added to the preliminary inventory and to obtain detailed
descriptions of each behavior. Additionally, these observations were supplemented with photographs
obtained during several years of observational study of this herd for other purposes, and with the
cumulative equid observational experience and study notes of the principal investigator with other
equid species. An initial draft was sent out to 18 equine behavior colleagues for review.
A total of 38 individual behaviors classi®ed into four distinct categories were included in the
ethogram. These included object play (14 entries), play sexual behavior (3 entries), locomotor play
(14 entries) and play ®ghting (7 entries). All of the behaviors catalogued from direct observation of
the herd were also found in the equid literature.
The resulting ethogram offers a practical tool as a ®eld guide or reference for quantitative research
and other studies of equid play behavior as well as for teaching of equid behavior.
# 2002 Elsevier Science B.V. All rights reserved.

Keywords: Equine; Pony; Zebra; Donkey; Przewalski horse; Play behavior; Ethogram

1. Background and purpose

Equid behavior has historically been a topic of great interest to animal keepers,
biologists, animal scientists, veterinarians, and ethologists. Numerous descriptive accounts
of equid behavior as well as a few systematic studies focusing on particular species or types
of behavior are available in the lay and scienti®c literature. In this body of published

*
Corresponding author. Tel.: ‡1-610-444-5800x2221; fax: ‡1-610-925-8124.
E-mail address: suemcd@vet.upenn.edu (S.M. McDonnell).

0168-1591/02/$ ± see front matter # 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 1 6 8 - 1 5 9 1 ( 0 2 ) 0 0 1 1 2 - 0
264 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

information, there are notable apparent inconsistencies in the terminology and interpreta-
tions of behavior.
An ethogram is a formal description of a species behavioral repertoire or a major
segment of it. It may be a complete list of all behaviors or it may focus on particular
functional classes of behaviors (Grier, 1984). To our knowledge, a complete ethogram for
equids has not been published. Our laboratory's long-term goal is to prepare a concise
comprehensive catalog-style equid ethogram, with the primary objective of establishing
standard nomenclature for equid behavior. To date, we published an ethogram of agonistic
intermale behavior of equid bachelor bands (McDonnell and Haviland, 1995). The
objective of the present work was to complete an ethogram of equid play behavior.
Play has been broadly characterized as activities appearing to have no immediate use or
function to the animal, involving a sense of pleasure and elements of surprise (McFarland,
1987). Across species, play behavior in large part appears to be a modi®ed form of serious
survival activities such as locomotor, aggressive, and reproductive behavior. In any species,
play is usually distinguished from serious forms of behavior by postures and expressions
denoting less serious ``intent'' (Schilder et al., 1984). In most species, play behavior occurs
both as a solitary activity and as a social interactive behavior. Play is believed to serve a
variety of adaptive functions, including enhancing general musculoskeletal and cardio-
vascular ®tness, practicing and honing speci®c survival skills, gaining familiarity with the
particular environment, or building social relationships and communication skills (Fagen,
1981; Bekoff and Byers, 1998).
Play behavior is a remarkably conspicuous feature of equid developmental behavior
(Fagen and George, 1977; Fagen, 1981). Study of play among free-running (wild, feral,
semi-feral) or domestic equids (mare and foal groups) has been limited mostly to relatively
informal mention, description, and/or simple classi®cations, with some exceptions. Fagen
and George (1977) quanti®ed the percentage of exercise attributable to play and non-play
activities of pony foals from birth to 6 weeks of age pastured in natural family bands.
Crowell-Davis (1983, 1986) reported the frequency, length, and type of play bouts in
domestic Welsh Pony foals at pasture with their dams in a mare and foal group, as well as
developmental changes, sex differences, and effect of environmental temperature on play
in young horses (1987). In addition, several authors included a considerable amount of play
in their descriptive studies of behavior and ecology of various equid populations. These
include Feist (1971), who studied feral horses in the Pryor Mountains along the Wyoming±
Montana border, USA; Tyler (1972), who described differences in play among domestic
foals of different ages and gender among a population of domestic horses and ponies kept
under semi-feral conditions for several months each year in the New Forest area of
England; Blakeslee (1974), who studied Appaloosa horses pastured under semi-natural
conditions in Idaho, USA; Moehlman (1974), who studied feral African wild asses of
Death Valley, CA, USA; Gardner (1983), who studied Grevy's zebra in Kenya; Schilder
et al. (1984) who studied plains zebra in a semi-preserve, differentiating facial expressions
of play from serious aggressive behavior; Penzhorn (1984), who studied Cape Mountain
zebra in South Africa; Keiper (1985), who studied feral ponies on the Assateague Island off
the coast of Maryland, USA; Hoffmann (1985), who studied a feral horse population off the
coast of North Carolina, USA; and Boyd et al. (1988), who studied time budgets in
Przewalski horses in a semi-preserve in Virginia, USA.
 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 265

2. Methods

2.1. General approach

The methods followed those used in the previously published intermale agonistic
ethogram for equids (McDonnell and Haviland, 1995). The general method included
review of existing equid behavior literature, original observations of equids, and incor-
poration of critical comment from equine behaviorists worldwide. A catalog format was
selected for simplicity and consolidation of information, as likely to be the most useful as a
practical ®eld or laboratory guide for equine behavior research.

2.2. Play behavior inventory from the equid literature

Literature (English-language) pertaining to semi-feral, feral, and domestic horses as

well as other captive or free-running equids was reviewed to generate an inventory of
behavioral elements and sequences displayed during play interactions.

2.3. Original observations

Our primary original observations were conducted within the context of a semi-feral
herd of Shetland-type ponies that resides at the University of Pennsylvania School of
Veterinary Medicine at New Bolton Center in Kennett Square, PA. This herd has been
maintained for approximately 8 years with minimal intervention. It includes multiple
harem groups with foals, yearlings and some young adult offspring, as well as bachelor and
young juvenile groups totaling 50±75 animals. With the objectives of developing a
complete inventory of play behavior, obtaining photo-documentation of each behavioral
entry, and preparing precise text descriptions and a line drawing depicting each entry, over
100 h of direct observation focused speci®cally on play behavior. These observations were
done during the late spring and summer of 2000. The observations were supplemented with
photographs obtained during several years of observational study of this herd for other
purposes, and with the cumulative equid observational experience and study notes of the
principal investigator with other equid species.

2.4. Collaborative review

A preliminary draft of this ethogram was circulated to 18 equine behavior researchers for
critical review; 12 responded with critical review (see `Acknowledgements' section).

3. Results

Table 1 represents our resulting ethogram. We classi®ed play into four categories,
similar to classi®cations proposed for mammals in general (Fagen, 1981). Object play
includes 14 entries, play sexual behavior includes 3 entries, locomotor play includes
266 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Table 1
The resulting ethogram comprising object play, sexual behavior, locomotor play and play fighting

1. Object play
Object play involves contact and manipulation of an object. The target object may be either inanimate such as
an environmental object, or an animate object such as the mane, tail or other body part of a herd mate or even of
an animal of another species. Object play has also been called object manipulation or manipulative play
(Crowell-Davis, 1986; Crowell-Davis et al., 1987; Waring, 1983). The following 14 entries represent commonly
observed equid object play behaviors.
Some of these behaviors, for example, circle and to and from, obviously involve locomotion. Although they
could easily be classified as locomotor play, they so clearly seem focused on an object, and often appear to be
stimulated by the object that we have included them with object play.

With jaws closed the upper lip is moved upward and downward against an object,
typically without dental contact of the object.
Comments: Nibbling of an object is typically one of the first play responses
associated with an investigative approach of the object. In many instances, nibbling
appears to be a means of moving an object on the ground (see Fig. 1).
Species: HorseÐsemi-feral ponies (Tyler, 1972); free-ranging Appaloosa horses
(Blakeslee, 1974); feral horses (Waring, 1983); feral ponies (Keiper, 1985); Welsh
Pony mare and foal groups (Crowell-Davis, 1983, 1986; Crowell-Davis et al.,
1987). DonkeyÐferal asses Equus africanus (Moehlman, 1998).

The tongue is extended through the teeth and border of the mouth, making contact with an
object. It is then retracted back into the mouth and chewing may follow. This sequence is
typically repeated.
Comments: Sniffing and/or licking an inanimate object may be as if to investigate the odor,
texture, shape, taste, and size of an object. Sniffing and licking of a herd mate sometimes
precedes and appears to initiate mutual grooming.
Species: HorseÐferal ponies (Keiper, 1985).

An object, whole or part depending on size, is taken into the mouth with upper and lower
lips and tongue then placed between the lips, incisors (front teeth) or molars. The head is
usually elevated once the object is in the mouth (see pick up). The animal may move
around with the object in the mouth (see carry).
Other names: Manipulate by mouth (Waring, 1983).
Comments: The animal may shake the object by tossing the head and neck while the
object is being mouthed. A dam's mane or tail is a common target of mouthing or
chewing.
Species: HorseÐfree-ranging Appaloosa horses (Blakeslee, 1974); domestic and feral
horses and ponies (Waring, 1983); Welsh Pony foals and mares (Crowell-Davis, 1983,
1986; Crowell-Davis et al., 1987). DonkeyÐferal asses Equus africanus (Moehlman,
1974).
 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 267

Table 1 (Continued )

An object is taken into the mouth with a side-to-side grinding motion of upper and
lower jaw. May include head tossing and/or forward movement of the tongue
through the front teeth ending with the object falling out of the mouth.
Comments: This behavior appears to be for the purpose of investigating the texture,
shape and/or size of the object, rather than for nourishment. The mother's mane and/
or tail are common target objects for chewing or mouthing by the foal. In horses and
ponies, all ages have been observed to chew inanimate objects or animate objects
(McDonnell and Poulin, unpublished observations).
Species: HorseÐsemi-feral ponies (Tyler, 1972); domestic and feral horses and
ponies (Waring, 1983).

An object is held between the lips, front teeth or molars. The head is elevated
with object in the mouth so that the object is lifted from the ground. The
height at which the object is lifted can vary from a few inches to several feet.
Other names: Lift (Waring, 1983).
Comments: See Figs. 2 and 3.
Species: HorseÐsemi-feral ponies (Tyler, 1972); domestic and feral horses
and ponies (Waring, 1983); Welsh Pony foals and mares (Crowell-Davis,
1983, 1986; Crowell-Davis et al., 1987). DonkeyÐferal asses Equus
africanus (Moehlman, 1974).

Following pick up, the object may be moved in a side-to-side, up-and-down,

or circular motion.
Other names: Swing head (Crowell-Davis, 1983; Crowell-Davis et al., 1987);
wave about (Crowell-Davis, 1983); scrape along ground (Moehlman, 1974)
Comments: See Figs. 2 and 3.
Species: HorseÐWelsh Pony foals and mares (Crowell-Davis, 1983, 1986;
Crowell-Davis et al., 1987). DonkeyÐferal asses Equus africanus (Moehlman,
1974).

Following pick up, the object is held between the jaws as the animal moves.
Other names: Drag (Moehlman, 1974; Crowell-Davis et al., 1987).
Species: HorseÐWelsh Pony foals and mares (Crowell-Davis, 1983, 1986;
Crowell-Davis et al., 1987).

Following pick up, the upper and lower jaws open, releasing the object
(drop) or throwing the nose upward as the object is released (toss). A toss
may alternately be effected without pick up by using the muzzle to flip an
object from the substrate into the air.
Comments: Crowell-Davis (1983) notes that Welsh Pony foals have been
observed tossing the mane and tail of their dam with their nose.
Species: HorseÐsemi-feral ponies (Tyler, 1972); domestic and feral horses
and ponies (Waring, 1983); Welsh Pony foals and mares (Crowell-Davis,
1983, 1986; Crowell-Davis et al., 1987). DonkeyÐferal asses Equus
africanus (Moehlman, 1974).
268 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Table 1 (Continued )

An object is held between the lips or front teeth, followed by a dragging

motion of the object with forward and back or side-to-side movement. The
head and neck or full body may move in any direction.
Other names: Drag (Crowell-Davis et al., 1987).
Species: HorseÐsemi-feral ponies (Tyler, 1972); domestic and feral horses
and ponies (Waring, 1983); Welsh Pony mares and foals at pasture (Crowell-
Davis, 1983; Crowell-Davis et al., 1987); feral ponies (Keiper, 1985).

With an object as an apparent target, one foreleg is lifted off the ground slightly,
extended quickly in a forward direction followed by a backward, toe-dragging move-
ment as if digging. The movement is typically repeated several times in succession
(McDonnell and Haviland, 1995). The foot may have direct contact with the object thus
moving the object. Alternatively, the foot may be slightly behind the object touching
the ground.
Comments: Investigative sequences and water play typically include pawing (see Figs. 4
and 5). Pawing an animate object also occurs as an apparent play initiation gesture (see
Fig. 6a and b).
Species: HorseÐsemi-feral ponies (Tyler, 1972); feral horses (Boyd, 1980); Welsh
Pony mares and foals (Crowell-Davis, 1983, 1986; Crowell-Davis et al., 1987);
domestic and feral horses (Waring, 1983); feral ponies (Keiper, 1985). DonkeyÐferal
asses Equus africanus (Moehlman, 1974). Przewalski horseÐferal mares and foals
(Boyd, 1980).

Standing at right angles to a herd mate target (usually the dam), with the butt
toward and often touching the abdomen of the target, weight is transferred to
the front legs as the hindlegs are raised in a hopping motion a few inches off the
ground toward the target. Typically, no extension of the hindlegs, as in kicking,
occurs.
Species: HorseÐsemi-feral ponies (Tyler, 1972); feral horses (Boyd, 1980).
ZebraÐGrevy's zebra in natural social groups (Gardner, 1983).

The equid moves, usually at a trot or gallop, away from an

object, such as the dam or a tree, and then returns to the
object at any gait.
Other names: Runs in opposite directions (Moehlman,
1974).
Comments: This behavioral sequence could also be con-
sidered as locomotor play, but is classified here as object
play because the action so obviously involves a landmark
object.
Species: HorseÐsemi-feral ponies (Tyler, 1972); pastured
domestic ponies in natural herd (Fagen and George, 1977);
feral horses (Boyd, 1980); domestic and feral horses and
ponies (Waring, 1983); feral ponies (Keiper, 1985).
DonkeyÐferal asses Equus africanus (Moehlman, 1974).
ZebraÐGrevy's zebra in natural social groups (Gardner,
1983).
 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 269

Table 1 (Continued )

The animal moves in a path, generally circular, around an

object so that the beginning and ending point are in the same
general vicinity. This behavior can be performed at any gait
and may be repeated.
Other names: Running in loops (Blakeslee, 1974).
Comments: The circling to be around an object. From our obser-
vations, this most frequently occurred in foals or adolescents,
and was rarely observed in mature animals. The focal animate
object was usually the dam (see Fig. 7a and b). The most
common inanimate focal object was a tall weed, or tree. May
also be classified as a locomotor behavior, but included here
with object play because of the distinct focus on an object.
Species: HorseÐsemi-feral ponies (Tyler, 1972); free-ranging
Appaloosa horses (Blakeslee, 1974); feral horses (Boyd, 1980);
Welsh Pony foals and mares (Crowell-Davis, 1983; Crowell-
Davis et al., 1987); feral ponies (Keiper, 1985). ZebraÐGrevy's
zebra in natural social groups (Gardner, 1983).

The animal raises its chest and forelegs so that one or both limbs rest across the
body of a herd mate, typically with lateral orientation.
Comments: See Fig. 8. The animal rearing may rotate around the partner's body so
that it ends up in the mounting position, part of sexual play. In our observations of
ponies, a considerable proportion of occurrences of resting rear are not within the
context of sexual play per se, but rather dispersed within bouts of locomotor play
and play fighting.
Species: HorseÐsemi-feral ponies (Tyler, 1972); free-ranging Appaloosa horses
(Blakeslee, 1974); domestic Shetland and Welsh Pony mares and foals at pasture
(Schoen et al., 1976); feral ponies (Keiper, 1985). DonkeyÐferal asses Equus
africanus (Moehlman, 1998).

2. Play sexual behavior

Sexual play is conspicuous and frequent in foals and young adolescents of both sexes, and among young and
adult bachelor stallions. The elements of the precopulatory and copulatory sequences may be out of order or
exaggerated from that of a mature adult in a breeding context. The following three entries represent commonly
observed play sexual behavior in equids.
The animal approaches and sniffs voided urine or feces,
performs the flehmen response, and covers the urine or feces
with urine and/or feces, and then again sniffs and performs
flehmen response, in the stylized postures typical of a mature
stallion but in less organized sequences.
Comments: Usually performed by foals and more frequently
by yearlings after elimination of a mature animal. Sequence
is often quite similar to adult form, except that it may not
proceed in the order typical of adult elimination marking
sequence and the elements may be interspersed among play
sequences of other types, for example, frolic.
Species: HorseÐferal horses (Feist, 1971); semi-feral ponies
(Tyler, 1972); free-ranging Appaloosa horses (Blakeslee,
1974); Shetland and Welsh Pony mares and foals at pasture
(Schoen et al., 1976); feral ponies (Keiper, 1985). Przewalski
horseÐferal mares and foals (Boyd, 1980).
270 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Table 1 (Continued )

One animal sniffs and/or nuzzles the head, shoulder, abdomen, flank,
inguinal, tail and/or genital areas of another in a similar, but more
playful, manner to that of an adult stallion investigating a mare before
copulation.
Comments: Foals of both sexes and particularly yearling colts often
tease a mare during or following a mature stallion's teasing or
breeding of the mare. The precopulatory sequence may or may not be
complete or in the order typical of a stallion. In play sequences, the
flehmen response occurs, but the tip of the nose is typically not raised
as high as a mature stallion (Keiper, 1985).
Species: HorseÐsemi-feral domestic ponies (Tyler, 1972); free-
ranging Appaloosa horses (Blakeslee, 1974); Shetland and Welsh
Pony mares and foals at pasture (Schoen et al., 1976); feral ponies
(Keiper, 1985). ZebraÐGrevy's zebra in natural social groups
(Gardner, 1983).

The animal raises its chest and forelegs onto the back of a herd mate
(same or opposite sex) as during copulation in mature adults. The mount
may be oriented from the side or rear.
Other names: Sexual mounting of mother (Blakeslee, 1974); sex without
coition (McFarland, 1987).
Comments: Usually performed by foals and immature young of both sexes
to their mother and other immature herd mates. It may occur with or
without either precopulatory (teasing, elimination marking) and copula-
tory (thrusting) behavioral elements. Sexual arousal as in an adult is
usually not apparent, although erection can be present. Lateral mount,
though similar in form to resting rear, by context and form appears
distinct from resting rear. Fig. 9 illustrates a typical play mounting
sequence. An abbreviated form of the mount may include only resting the
chin and head on the hindquarters of the target animal (as if about to
mount).
Species: HorseÐferal horses (Feist, 1971); semi-feral ponies (Tyler,
1972); free-ranging Appaloosa horses (Blakeslee, 1974); Shetland and
Welsh Pony mares and foals at pasture (Schoen et al., 1976); feral horses
(Boyd, 1980); domestic horse (Fagen, 1981); domestic and feral horses
and ponies (Waring, 1983); Welsh Pony foals and mare groups (Crowell-
Davis, 1983, 1986; Crowell-Davis et al., 1987); feral ponies (Keiper,
1985). DonkeyÐferal asses Equus africanus (Moehlman, 1998). ZebraÐ
Grevy's zebra in natural social groups (Gardner, 1983); Cape Mountain
zebra Equus zebra zebra (Penzhorn, 1984).
 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 271

Table 1 (Continued )

3. Locomotor play
Locomotor play involves any play behavior response or sequence that is performed while in motion at any
gait. Circling and to and from were included in object play because the action appears to be directed toward or
around an object/s, but each can also be considered locomotor play.
The following seven entries represent common locomotor play behaviors in equids. Certainly, there are many
more movements that could be considered as distinct behaviors. For example, play also includes a variety of
head movements, including head toss, head twist, and head shake (Fagen and George, 1977).
With the exception of chase, which is by definition social, each of these locomotor play behaviors can occur
either in a solitary or a social context.
Fore- and hindlegs simultaneously propel off the ground along with apparently
exuberant, random bucking, head shaking and body twists. May take off from a
stationary position to an instantaneous gallop.
Other names: Cavorting (Fagen, 1981); gamboling (McFarland, 1987; Fagen,
1981); capering (Fagen, 1981); prop (Fagen and George, 1977).
Comments: May interrupt an episode of running.
Species: HorseÐdomestic pony foals pastured in a natural herd (Fagen and
George, 1977); domestic horse foals at pasture with dams (Fagen, 1981); domestic
and feral horses and ponies (Waring, 1983); Welsh Pony foals and mares (Crowell-
Davis, 1983; Crowell-Davis et al., 1987). DonkeyÐferal asses Equus africanus
(Moehlman, 1998). ZebraÐGrevy's zebra in natural social groups (Gardner,
1983).

With no apparent destination to reach or threat to escape, the animal moves at the
canter or gallop in a seemingly spontaneous burst of motion.
Other names: Galloping (Keiper, 1985; Tyler, 1972); exuberant galloping
(Waring, 1983).
Comments: Fagen and George (1977) reported that greater than two-thirds of all
running in pastured pony foals from birth to 6 weeks was in a play context.
Fig. 10 illustrates solitary running in a young foal.
Species: HorseÐferal horses (Feist, 1971); semi-feral ponies (Tyler, 1972); free-
ranging Appaloosa horses (Blakeslee, 1974); Shetland and Welsh Pony mares
and foals at pasture (Schoen et al., 1976); domestic pony foals pastured in a
natural herd (Fagen and George, 1977); domestic horse foals with mares (Fagen,
1981); domestic and feral horses and ponies (Waring, 1983); Welsh Pony mare
and foal groups (Crowell-Davis, 1983; Crowell-Davis et al., 1987); feral ponies
(Keiper, 1985); feral horses (Berger, 1986). DonkeyÐferal asses Equus
africanus (Moehlman, 1998). Przewalski horseÐferal mares and foals (Boyd,
1980). ZebraÐGrevy's zebra in natural social groups (Gardner, 1983).
272 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Table 1 (Continued )

At the trot, canter or gallop, an animal is pursued, with the apparent

effort to catch up to and overtake it.
Other names: Charge (Keiper, 1985); race (Penzhorn, 1984).
Comments: The animal often attempts to nip at or bump (similar to
push at speed) the pursued play partner while chasing. The participants
may reverse roles of chasing and being pursued. Play chasing can
occur as what appears to be simple locomotor ``games'' or ``tag'' or
can occur within the context of play fighting. Fig. 11 illustrates a group
chase. Chasing, particularly in play among bachelors, can also appear
to be herding similar to that of a harem stallion herding and driving his
mares (McDonnell and Haviland, 1995).
Species: HorseÐferal horses (Feist, 1971); semi-feral ponies (Tyler,
1972); free-ranging Appaloosa horses (Blakeslee, 1974); Shetland and
Welsh Pony mares and foals at pasture (Schoen et al., 1976); domestic
ponies pastured in a natural herd (Fagen and George, 1977); feral
horses (Boyd, 1980); domestic and feral horses and ponies (Waring,
1983); immature male feral horses (Hoffmann, 1985); feral ponies
(Keiper, 1985); feral horses of the Great Basin (Berger, 1986). ZebraÐ
Cape Mountain zebra Equus zebra zebra (Penzhorn, 1984).

With the head and neck lowered and the weight shifted to the forelegs,
both hindlegs lift off the ground and extend backwards simultaneously,
often repeatedly in rapid succession.
Other names: Kicking of hindlegs into the air (Feist, 1971; Keiper, 1986);
kicking up (Blakeslee, 1974); kick out (Tyler, 1972; Fagen, 1981); rear
kicking (Fagen, 1981).
Species: HorseÐferal horses (Feist, 1971); semi-feral ponies (Tyler,
1972); free-ranging Appaloosa horses (Blakeslee, 1974); Shetland and
Welsh Pony mares and foals at pasture (Schoen et al., 1976); domestic
ponies pastured in a natural herd (Fagen and George, 1977); feral horses
(Boyd, 1980); domestic horse foals at pasture with dams (Fagen, 1981);
domestic and feral horses and ponies (Waring, 1983); Welsh Pony mare
and foal groups (Crowell-Davis, 1983, 1986; Crowell-Davis et al., 1987);
feral ponies (Keiper, 1985); feral horses (Berger, 1986). DonkeyÐferal
asses Equus africanus (Moehlman, 1998). ZebraÐGrevy's zebra in
natural social groups (Gardner, 1983).

With mostly hindlimb propulsion, the animal suddenlyA Ã moves forward with the
forelegs leaving the ground first followed by the hindlegs. Can either appear to
be jumping over a target obstacle or not.
Species: HorseÐsemi-feral ponies (Tyler, 1972); domestic horse foals at pasture
with dams (Fagen, 1981); domestic and feral horses and ponies (Waring, 1983).
DonkeyÐferal asses Equus africanus (Moehlman, 1998).
 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 273

Table 1 (Continued )

In a combination of jump and frolic, the animal propels itself

off the ground with its forelegs and hindlegs over an object,
away from an object, or toward an object.
Comments: In the context of foal locomotor play and play
as an adult, leap can sometimes resemble frolic, however,
the leap action is directed toward or against another animal
(Crowell-Davis, 1983, 1986; Fagen, 1981; McFarland,
1987).
Species: HorseÐdomestic horse foals (Fagen, 1981); Welsh
Pony foals and mares (Crowell-Davis, 1983, 1986; Crowell-
Davis et al., 1987). DonkeyÐferal asses Equus africanus
(Moehlman, 1998).

The animal walks or trots forward with neck arched, ears forward, tail elevated,
and exaggerated knee action. The head and neck may bob up-and-down while in
motion, and a snorting sound may be emitted with each stride.
Comments: Often occurs at the end of a play bout.
Species: HorseÐsemi-feral ponies (Tyler, 1972); free-ranging Appaloosa horses
(Blakeslee, 1974); feral horses (Boyd, 1980); Welsh Pony mare and foal groups
(Crowell-Davis, 1983, 1986; Crowell-Davis et al., 1987); feral ponies (Keiper,
1985).

4. Play fighting
Play fighting involves behavioral elements and sequences similar to serious adult fighting behavior, but with
more of a sporting character than serious fighting. In contrast to serious fights, the cohorts appear to alternate
offensive and defensive roles, spar on as if to ``keep the game going,'' and stop short of injury.
Initiation of play fighting has been observed to follow the head threat tossing motion in which the head is
rapidly flipped up-and-down, either while the animal is standing still or moving. Play fighting has been
described by Berger (1986), Keiper (1985), Moehlman (1998) and Schilder et al. (1984). Play fighting has also
been termed interactive (contact or combat) play (Crowell-Davis, 1983, 1986; Crowell-Davis et al., 1987) and
aggressive play (Tyler, 1972). The following 14 entries represent common play fighting behaviors in equids.
For the behaviors involving mouth aggression, we distinguished three types of action. We used the term nip to
refer to a slight opening of the jaw to take and quickly release small pieces of skin or flesh between the teeth;
bite to refer to a wider opening of the jaws and teeth to take and quickly release a larger piece of flesh and skin
between the teeth; and grasp to refer to an extended clamping hold of a larger piece of flesh and skin, or a limb,
between widely opened jaws.
Most play fighting occurs in a social context. However, we have observed certain elements such as stamp and
rear within solitary play bouts of young foals.
274 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Table 1 (Continued )

Nip: the jaws and teeth are opened and closed slightly taking a
small piece of hair or flesh of a cohort between the teeth. Bite: the
jaws and teeth are opened widely, closed and quickly released on a
large piece of flesh and skin of a cohort between the teeth. The ears
are upright and lips may be retracted.
Other names: Neck biting (Gardner, 1983); face nip (Blakeslee, 1974).
Comments: Fig. 12 illustrates nipping the head. Nipping and biting
may occur in various areas of the body, including in addition to the
forebody, the stifle and flank. Fig. 13 depicts a recumbent foal
biting the chest of a cohort that has just nudged to invite play.
Species: HorseÐsemi-feral ponies (Tyler, 1972); Shetland and
Welsh Pony mares and foals at pasture (Schoen et al., 1976); free-
ranging Appaloosa horses (Blakeslee, 1974); domestic and feral
horses and ponies (Waring, 1983); feral ponies (Keiper, 1985).
DonkeyÐferal asses Equus africanus (Moehlman, 1998). ZebraÐ
Grevy's zebra in natural social groups (Gardner, 1983); Cape
Mountain zebra Equus zebra zebra (Penzhorn, 1984).

With the jaws open and clamped, the mane and neck of a cohort are held at
the crest and sometimes moved back and forth.
Other names: Grip neck or mane (Keiper, 1985); holding mane (Feist,
1971); neck grip (Boyd, 1980; Tyler, 1972); holding crest (Crowell-Davis,
1983); grasp (McDonnell and Haviland, 1995); mane grip (Schoen et al.,
1976); biting (Moehlman, 1974).
Comments: May precede (as if to initiate) or follow (as if initiated by)
mutual grooming (McDonnell, unpublished observations), ponies.
Species: HorseÐferal horses (Feist, 1971); semi-feral ponies (Tyler, 1972);
Shetland and Welsh Pony mare and foals at pasture (Schoen et al., 1976);
feral horses (Boyd, 1980); domestic horse mare and foal groups (Fagen,
1981); Welsh Pony mare and foal groups (Crowell-Davis, 1983; Crowell-
Davis et al., 1987); immature male feral horses (Hoffmann, 1985); feral
ponies (Keiper, 1985); feral horses (Berger, 1986); semi-feral bachelor
bands (McDonnell and Haviland, 1995). DonkeyÐferal asses Equus
africanus (Moehlman, 1974, 1998). ZebraÐCape Mountain zebra
Equus zebra zebra (Penzhorn, 1984); plains zebra (Berger, 1986).

Sparring with the head and neck. One or both partners may remain
standing, drop to one or both knees, or raise the forelegs during a bout of
neck wresting. This activity may including pushing and slamming with
the shoulder against the shoulder or abdomen of the partner.
Other names: Neck fencing (Berger, 1986); wrestle (Hoffmann, 1985).
Species: HorseÐferal horses (Feist, 1971); semi-feral ponies (Tyler,
1972); free-ranging Appaloosa horses (Blakeslee, 1974); Shetland and
Welsh Pony mares and foals at pasture (Schoen et al., 1976); feral horses
(Boyd, 1980); domestic horse groups (Fagen, 1981); immature male feral
horses (Hoffmann, 1985); feral ponies (Keiper, 1985); feral horses
(Berger, 1986). DonkeyÐferal asses Equus africanus (Moehlman, 1974,
1998). ZebraÐCape Mountain zebra Equus zebra zebra (Penzhorn,
1984).
 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 275

Table 1 (Continued )

Nip: the jaws and teeth are opened and quickly closed and then released on a
small amount of hair or skin of a foreleg of the target animal. Bite: the jaws
and teeth are opened and closed slightly taking a large portion of flesh of the
target animal between the teeth. Grasp: the jaws and teeth are opened and
clamped in an extended hold of the limb of the target animal between the
teeth and jaws.
Comments: Typically causes the recipient to buckle at the knees and partially
drop to the ground with the front end, in an attempt to evade continued
contact (See Fig. 14). As in that photograph the two participants may
continue nipping each other from the ground position. In bachelor stallions,
this ground position has been called kneeling (McDonnell and Haviland,
1994). Keiper (1985) noted that this has also been called the ``turtle
posture.''
Species: HorseÐsemi-feral ponies (Tyler, 1972); free-ranging Appaloosa
horses (Blakeslee, 1974); Shetland and Welsh Pony mares and foals at
pasture (Schoen et al., 1976); domestic and feral horses and ponies (Waring,
1983); feral ponies (Keiper, 1985); feral horses of the Great Basin (Berger,
1986); Welsh Pony foal and mare groups (Crowell-Davis, 1986). ZebraÐ
Cape Mountain zebra Equus zebra zebra (Penzhorn, 1984); plains zebra
(Berger, 1986). Onagers (Berger, 1986).

Nip: the jaws and teeth are opened and quickly closed and then released on
a small amount of hair or skin of a hindleg of the opposing participant.
Bite: the jaws and teeth are opened and closed slightly taking a large
portion of flesh of the target animal between the teeth. Grasp: the jaws and
teeth are opened and clamped in an extended hold of the limb of the play
partner between the teeth and jaws.
Comments: Fig. 15 illustrates a young foal grasping the hindleg of a
yearling. Mutual hindleg nipping, biting, and grasping typically lead to
circling of two participants in an apparent attempt of each to avoid being
bitten while continuing to bite (see Fig. 16).
Species: HorseÐsemi-feral ponies (Tyler, 1972); free-ranging Appaloosa
horses (Blakeslee, 1974); Shetland and Welsh Pony mares and foals at
pasture (Schoen et al., 1976); domestic and feral horses and ponies
(Waring, 1983); feral ponies (Keiper, 1985). ZebraÐCape Mountain zebra
Equus zebra zebra (Penzhorn, 1984).

The jaws and teeth are opened and closed and then quickly released taking a small
(nip) or large (bite) piece of flesh on the posterior section of the rump.
Comments: May be followed with a buck from the recipient. Schoen et al. (1976)
note that tail biting is a component of play fighting.
Species: HorseÐsemi-feral ponies (Tyler, 1972); free-ranging Appaloosa horses
(Blakeslee, 1974); domestic and feral horses and ponies (Waring, 1983); feral ponies
(Keiper, 1985). ZebraÐCape Mountain zebra Equus zebra zebra (Penzhorn, 1984).
276 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Table 1 (Continued )

The head, neck, shoulders, chest, body or rump is pressed against the play
partner in an apparent attempt to displace the other (McDonnell and
Haviland, 1995). Sometimes in a single episode, two participants alternate
roles of pushing and being pushed.
Other names: Bump against (Gardner, 1983); push and bunt (Crowell-
Davis, 1986).
Comments: Fig. 17 illustrates a young foal pushing an older foal.
Species: HorseÐferal horses (Feist, 1971); domestic and feral horses and
ponies (Waring, 1983); Welsh Pony mare and foals (Crowell-Davis, 1983,
1986; Crowell-Davis et al., 1987); feral horses (Berger, 1986). ZebraÐ
Grevy's zebra in natural social groups (Gardner, 1983).

One foreleg is raised and lowered, sharply and firmly striking the
ground, sometimes repeated in a quick burst typically within the context
of a play fight sequence. May serve to emit an acoustical signal.
Other names: Strike (Boyd, 1980; Crowell-Davis, 1983; Waring, 1983);
front hoof beating (Hoffmann, 1985); paw (Schoen et al., 1976); stomp
(McDonnell and Haviland, 1995).
Species: HorseÐShetland and Welsh mare and foals at pasture (Schoen
et al., 1976); feral horses (Boyd, 1980); domestic and feral horses and
ponies (Waring, 1983); Welsh Pony mare and foal groups (Crowell-
Davis, 1983); immature male groups of feral horses (Hoffmann, 1985).

The forequarters are raised high while the hindlegs remain on the
ground, resulting in a near-vertical position (McDonnell and Haviland,
1995).
Comments: Fig. 18 illustrates a rear. May result in the resting rear
position or the partial mount or dancing or boxing (McDonnell and
Haviland, 1995; see Fig. 19). Commonly interpreted as related to
establishment of dominance during a play fight sequence.
Species: HorseÐdomestic ponies pastured in a natural herd (Fagen and
George, 1977); feral horses (Berger, 1986); free-ranging Appaloosa
horses (Blakeslee, 1974); feral horses (Boyd, 1980); semi-feral ponies
(Keiper, 1985); Welsh Pony mare and foal groups (Crowell-Davis, 1983,
1986; Crowell-Davis et al., 1987); domestic horse (Fagen, 1981);
Shetland and Welsh mare and foals at pasture (Schoen et al., 1976);
semi-feral ponies (Tyler, 1972); domestic and feral horses and ponies
(Waring, 1983). DonkeyÐferal asses Equus africanus (Moehlman, 1998).
ZebraÐCape Mountain zebra Equus zebra zebra (Penzhorn, 1984).

Usually with ears back and the rump turned toward a herd mate, an animal
raises one leg as if aiming to kick, often simultaneously backing toward the
target.
Other names: Kick threat (McDonnell and Haviland, 1995).
Comments: May occur in the context of a play fight sequence, sometimes
appearing to signal the termination of the play fight (Keiper, 1986).
Species: HorseÐfree-ranging Appaloosa horses (Blakeslee, 1974); domestic
and feral horses and ponies (Waring, 1983). DonkeyÐferal asses Equus
africanus (Moehlman, 1974). ZebraÐCape Mountain zebra Equus zebra
zebra (Penzhorn, 1984).
 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 277

Table 1 (Continued )

One hindleg is lifted off the ground and extended backwards usually
toward the play partner, rarely with sufficient reach or force to touch or
cause injury to the recipient. May be repeated in succession.
Other names: Hindkicking (Boyd, 1980); kick out (Gardner, 1983; Tyler,
1972).
Comments: A double hindleg kick seems more forceful than play, and
often seems to terminate the play fight sequence, so may not represent
play.
Species: HorseÐsemi-feral ponies (Tyler, 1972); feral horses (Boyd,
1980); domestic and feral horses and ponies (Waring, 1983); Welsh Pony
mare and foal groups (Crowell-Davis et al., 1987). DonkeyÐferal asses
Equus africanus (Moehlman, 1998). ZebraÐGrevy's zebra in natural
social groups (Gardner, 1983); Cape Mountain zebra Equus zebra zebra
(Penzhorn, 1984).

As two cohorts approach, one stops abruptly, reversing direction of

the forebody, typically withdrawing the head and neck (and
sometimes forebody) in a sweeping dorsolateral motion while the
hindbody remains or pivots in place.
Other names: Swerving (Waring, 1983); quick stop (Moehlman,
1974).
Species: HorseÐsemi-feral pony bachelor bands (McDonnell and
Haviland, 1995). DonkeyÐferal asses Equus africanus (Moehlman,
1974).

During social locomotor play and play fighting, contact is avoided by

propelling the fore, rear, or entire body off the ground away from the offensive
gesture.

Comments: In a given sequence play opponents often alternate evading and

attacking roles.
Species: HorseÐfree-ranging Appaloosa horses (Blakeslee, 1974); feral ponies
(Keiper, 1985).

During social locomotor play and play fighting, contact is avoided by

turning away from the offensive gesture in a quick, sharp motion
pivoting around one hindleg.
Other names: Whirling to avoid being bitten (Boyd, 1980).
Species: HorseÐferal horses (Boyd, 1980); Welsh Pony mare and
foal groups (Crowell-Davis, 1983; Crowell-Davis et al., 1987);
domestic and feral horses and ponies (Waring, 1983).
278 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Fig. 1. Foal nibble novel paper.

7 entries, and play ®ghting includes 14 entries. Within these categories comments were
included on social and solitary forms (Figs. 1±21).
The references to literature were chosen as readily available sources of further
information. We did not attempt to provide an exhaustive inventory of all citations nor
did we attempt to ®nd the earliest description of the behavior. Photographs illustrating
context and interactive sequences were appended.

Fig. 2. Solitary object playÐshake novel object (paper bag).

S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 279

Fig. 3. Social object playÐshake novel object (paper bag).

Fig. 4. Paw in water.

Fig. 5. Solitary object playÐpaw novel object (tarp).

280 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Fig. 6. Play initiation sequence: (a) nip neck, (b) paw back of recumbent, (c) paw back of rising participant,
followed by (d) resting rear, and (e) foreleg grasp of standing participant.

4. Discussion

Included in our ®nal list of equid play behaviors were what appeared in our original
observations and in reports from the literature to be the most common play behaviors of
equids. Some of our decisions concerning inclusion or exclusion from the ethogram may be
questionable. Certainly, there could easily have been many more entries, further delineating
S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 281

Fig. 7. Foal circle dam.

Fig. 8. Resting rear.

282 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Fig. 9. Male foal mount another male foal (a) from side, (b) from rear, and (c) from rear (note commencing
erection).

speci®c elements within play sequences. For example, elimination marking sequence could
have been presented as several separate entries, i.e. ¯ehmen, urinate on, defectate on, sniff
feces, sniff urine. Another example would be in play ®ghting sequences. Elements such as
kneeling, levade, lunge, and various threat gestures such as head toss have in other works
been considered distinct ethogram entries.
Similarly, decisions were not always clear as to whether particular behaviors associated
with play should be considered as elements of play or as initiators or terminators of play.
For example, social play bouts often are preceded by a mutual head toss or alert posture
(McDonnell and Haviland, 1995). It is not clear to us whether these represent play or play
initiation gestures. A related dif®culty was deciding whether or not certain behaviors had a
play form or were always serious behavior in young foals. For example, some authors have
considered mutual grooming, self-grooming, or rolling as play behavior (McGreevy, 1996;
Tyler, 1972). It is dif®cult to know whether or not these behaviors serve a serious grooming
purpose as in adults or represent play forms of the behavior. In our observations of ponies
 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 283

Fig. 10. Solitary run.

and horses, for each of these examples, the behavior occurred in complete form and
sequence with seemingly serious intent similar to that of adults. So, for example, in mutual
grooming the nips were as complete as those among adults, even in week-old foals. One
reviewer (S. Ralston, personal communication) indicated that she had observed rolling in
domestic foals that she interpreted as play rolling in response to novel substrate or a fresh
puddle.
An example where the distinction was not as clear concerns grazing or drinking
behavior. Before foals begin ingesting grass or drinking water, they all appear to go
through a brief developmental stage during which they seem to mimic the grazing or
drinking behavior of adults in a playful investigative manner, mouthing and chewing, but
not ingesting (Blakeslee, 1974). In our observations of pony and horse foals, this phase of
playful grazing and drinking is typically very brief, often less than a day. Once a foal begins

Fig. 11. Group chase.

284 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Fig. 12. Head nip.

Fig. 13. Bite chest during play initiation interaction.

 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 285

Fig. 14. Foreleg bite and grasp.

ingesting the grass or drinking water, the playful, investigative form is not seen again. The
foal may occasionally play with an unusual piece of vegetation, for example, a tall weed
encountered while grazing. It was with that logic that we did not include in this ethogram
behaviors for which the play form was very brief and apparently developmentally
transitional.
An interesting feature of play, particularly in young foals, is that typically sequences do
not include the vocalization included in the serious adult form of the behavior with the
exception of frolic. For example, play ®ghting does not include the grunts and squeals so

Fig. 15. Hindleg grasp.

286 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Fig. 16. Older and younger foals circling as each nips or bites the other's hindleg.

typical of serious ®ghting. As reported by Fagen and George (1977), vocalizations of

young foals (approximately 3 months of age and younger) are almost exclusively limited to
calls and answers to the dam. During frolic, pony and horse foals may snort and emit
audible squeals.

Fig. 17. Push.

 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 287

Fig. 18. Rear.

In our laboratory work over the years, we have had opportunities to observe young
equids in natural family groups, in domestic mare and foal pairs, and in pasture herds of
mare±foal pairs. Those observations suggest that there may be some differences in the
quantity and quality of play dependent on the social environment in which a foal develops.
First, among equids living in natural social harem and bachelor group organization, there
seems to be considerably more play in foals, yearlings, young adults, and some mature

Fig. 19. Rear with ``dancing'' (within a play fighting sequence of young adult male ponies).
288 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Fig. 20. Nose-to-nose investigation preceding play bout.

adults than in mare and foal pairs or herds. Several factors may account for these perceived
differences, including mixed ages, a preponderance of young animals, presence of bachelor
and harem stallions, and general social facilitation. For example, young foals and yearlings
appear to mutually stimulate play, seemingly increasing the amount of locomotor social
play in the younger foals, and likely increasing the distance that young foals play away
from the dam. Also, harem stallions engage in play with the foals and yearlings, seemingly
increasing the amount of play compared to that seen in mare±foal only groups. Similarly, in
natural equid populations, the bachelor stallions play ®ght and chase with yearlings and 2-
year-old, possibly further increasing the amount of social locomotor play and play ®ghting.
In a study of ten Shetland and Welsh Ponies kept at pasture in a fairly natural social band
(Fagen and George, 1977) play accounted for most locomotor exercise of foals from birth
to 6 weeks of age (from 54 to 84% of locomotor steps and from 85 to 100% of turns). In

Fig. 21. Mutual grooming in this instance preceding social play bout.
 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290 289

comparing species, it has similarly been suggested that social organization likely affects
the amount and type of play (Fagen, 1981). Among the territorial zebra and donkeys, where
mares and their offspring travel together unassociated with others, the frequency of social
play among foals is rare compared to that of the harem equids (Moehlman, 1974; Gardner,
1983).
Our observations were similar to reports in the literature (Blakeslee, 1974; Moehlman,
1974) that play frequently appears to be stimulated by novel conditions or stimuli, and so is
often interspersed with investigation and exploration. This seems particularly the case for
young foals. Novel situations such as changes in weather, for example, wind or precipita-
tion, or encounter of novel objects, or movement of the herd to areas not yet traveled by the
neonate are associated with an apparent increased frequency and duration of play (see
Figs. 1±3 and 5).
There are several distinct behaviors that appear to invite or initiate play. These include
nose-to-nose approach, nudging, nipping, tossing the head, and pawing at the prospective
play partner. Fig. 20 illustrates the common nose-to-nose investigation greeting commonly
preceding play bouts. Fig. 6 illustrates a commonly observed play initiation sequence
involving a standing foal pestering a recumbent foal until it rises and joins in the play. As
noted by others (Schoen et al., 1976; Crowell-Davis et al., 1987) mutual grooming is
sometimes associated with play, either preceding or following a play bout (see Fig. 21).
There also appear to be certain actions associated with termination of a play bout.
Pinning the ears back, forceful biting, turning the rump toward the other animal and kicking
or striking with apparent serious intent often abruptly terminate a play bout. While these
actions are very similar to those occurring in play, when observed in the context of
terminating a play bout there are distinct differences in the apparent force and intent.
Another common ending event of social play bouts include one or more participants
prancing off.
It is worth noting that, while play is likely most frequent and conspicuous in foals,
yearlings, and bachelors, play also occurs in mature adults. Among adults, it appears most
common in bachelor stallions. Within harem groups, it appears to be more common in
harem stallions than in harem mares.

Acknowledgements

This is a Dorothy Russell Havemeyer Foundation project. Melissa Owen and Beth
Stephano assisted with ®eld photography. Line drawings were prepared by Erlene
Michener. The following colleagues provided critical review of an earlier draft: Carys
Hughes, Lee Boyd, Daniel Mills, Barbara Simpson, Sarah Ralston, Malgorzata Pozor,
Nancy Diehl, Machteld van Dierendonck, Corrina Snook, Jodi Drake, Jocelyn Kish, and
Sarah Strand.

References

Bekoff, M., Byers, J.A., 1998. Animal Play Evolutionary, Comparative, and Ecological Perspectives. Cambridge
University Press, Cambridge, UK.
290 S.M. McDonnell, A. Poulin / Applied Animal Behaviour Science 78 (2002) 263±290

Berger, J., 1986. Wild Horses of the Great Basin Social Competition and Population Size. The University of
Chicago Press, Chicago.
Blakeslee, J.K., 1974. Mother±young relationships and related behavior among free-ranging Appaloosa horses.
Master's thesis, Idaho State University, Pocatello.
Boyd, L.E., 1980. The natality, foal survivorship, and mare±foal behavior of feral horses in Wyoming's Red
Desert. Master's thesis, University of Wyoming, Laramie.
Boyd, L.E., Carbanaro, D.A., Houpt, K.A., 1988. The 24 hour time budget of Przewalski horses. Appl. Anim.
Behav. Sci. 21, 5±17.
Crowell-Davis, S.L., 1983. The behavior of Welsh Pony foals and mares. Ph.D. thesis, Cornell University,
Ithaca, NY.
Crowell-Davis, S.L., 1986. Developmental behavior. In: Crowell-Davis, S.L., Houpt, K.A. (Eds.), The
Veterinary Clinics of North America, Equine Behavior. Saunders, Philadelphia, pp. 573±590.
Crowell-Davis, S.L., Houpt, K.A., Kane, L., 1987. Play development in Welsh Pony (Equus caballus) foals.
Appl. Anim. Behav. Sci. 18, 119±131.
Fagen, R.M., 1981. Animal Play Behavior. Oxford University Press, New York.
Fagen, R.M., George, T.K., 1977. Play behavior and exercise in young ponies (Equus caballus L.). Behav. Ecol.
Sociobiol. 2, 267±269.
Feist, J.D., 1971. Behavior of feral horses in the Pryor Mountain Wild Horse Range. Master's thesis, University
of Michigan, Ann Arbor.
Gardner, C.D., 1983. Grevy's zebra of Samburu Kenya: mothers and foals project in wildlife ecology. Master's
thesis, Yale University, New Haven, CT.
Grier, J.W., 1984. Biology of Animal Behavior. Times Mirror/Mosby College Publishing, St. Louis, MO.
Hoffmann, R., 1985. On the development of the social behavior in immature males of a feral horse population
(Equus przewalskii f. caballus). Z. Saugetierkd. 50, 302±314.
Keiper, R., 1985. The Assateague Ponies. Tidewater Publishers, Centreville, MD.
McDonnell, S.M., Haviland, J.C.S., 1995. Agnostic ethogram of the equid bachelor band. Appl. Anim. Behav.
Sci. 43, 147±188.
McGreevy, P., 1996. Why does my horse. . .? Trafalger Square Publications, North Pomfret, VT, USA.
McFarland, D., 1987. The Oxford Companion to Animal Behavior. Oxford University Press, New York.
Moehlman, P.D., 1974. Behavior and ecology of feral asses (Equus asinus). Ph.D. dissertation, University of
Wisconsin, Madison.
Moehlman, P.D., 1998. Behavioral patterns and communication in feral asses (Equus africanus). Appl. Anim.
Behav. Sci. 60, 125±169.
Penzhorn, B.L., 1984. A long-term study of social organization and behavior of Cape Mountain zebras Equus
zebra zebra. Z. Tierpsychol. 64, 97±146.
Schilder, M.B.H., van Hoof, J.A., van Geer-Plesman, C.J., Wensing, J.B., 1984. A quantitative analysis of facial
expression in the plains zebra. Z. Tierpsychol. 66, 11±32.
Schoen, A.M.S., Banks, E.M., Curtis, S.E., 1976. Behavior of young Shetland and Welsh Ponies (Equus
caballus). Biol. Behav. 1, 192±216.
Tyler, S.J., 1972. Behavior and social organization of the semi-feral ponies. Anim. Behav. Monogr. 5.
Waring, G.H., 1983. Horse Behavior: The Behavioral Traits and Applications of Domestic and Wild Horses,
Including Ponies. Noyes Publications, Park Ridge, NJ.